59 The Neural Correlates of Language Production

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59
The Neural Correlates
of Language Production
PETER INDEFREY A N D WILLEM J. M. LEVELT
ABSTRACT This chapter reviews the findings of 58 word production experiments using different tasks and neuroimaging
techniques. The reported cerebral activation sites are coded in
a common anatomic reference system. Based on a functional
model of language production, the different word production
tasks are analyzed in terms of their processing components. This
approach allows a distinction between the core process of word
production and preceding task-specific processes (lead-in processes) such as visual or auditory stimulus recognition. The core
process of word production is subserved by a left-lateralized
perisylvian/thalamic language production network. Within this
network there seems to be functional specialization for the processing stages of word production. In addition, this chapter includes a discussion of the available evidence on syntactic
production, self-monitoring, and the time course of word production.
In reading the neuroscience literature on language production, one might infer that producing language simply
means producing words. Neuroimaging studies of language production typically require subjects to generate
(silently) words in response to other words (as in verb generation)—words of a particular semantic category, names
of depicted objects, words beginning with a particular
phoneme (or letter), and the like. Such studies have provided a wealth of information on the neurophysiology of
lexical access, but they should not obscure our perspective on the larger speech production process. Speaking is,
after all, our most complex cognitive-motor skill, designed by evolution to support communication in large
clans of homo sapiens. A vast network of brain structures,
both cortical and subcortical, contributes to the highspeed generation of utterances in never-identical communicative settings. It also generates the ever-babbling internal speech, speech whose representational functions are
still fallow research territory.
In this chapter, therefore, we begin with a summary
outline of the functional organization of speaking, laying
out the processing components involved, including
grammatical encoding, phonological encoding, and selfmonitoring. These components then offer a structure for
the subsequent review of neuroimaging studies, most of
them word production studies.
The functional organization of language production
The interactive generation of utterances in conversation,
the evolutionary basic setting for language use, involves
a multicomponent processing system. It can map communicative intentions onto articulatory gestures, which
in turn produce the auditory signals from which the interlocutor can derive or recognize these intentions. Figure 59.1 diagrams the major processing components
involved (roughly as defined in Levelt, 1989). Although
the modeling of component processes and their interaction still differs substantially among theories of language
production (see, in particular, the B B S commentaries to
Levelt, Roelofs, and Meyer, 1999), there is reasonable
consensus about the major components involved in the
generation of speech.
It makes both functional and neuropsychological
sense to partition these components as follows. There
is, on the one hand, a rhetorical/semantic/syntactic system. It decides on the communicatively effective information to express, puts it in terms of linguistically
expressible conceptual structures (“messages”), whereupon these messages trigger the generation of ordered
lexicosyntactic structures (“surface structures”). On the
other hand, there is also a phonological/phonetic system whose aim it is to generate the appropriate articulatory shape for these surface structures. Both systems
have access to a huge mental lexicon. The rhetorical/
semantic/syntactic system has, in addition, access to
communicatively relevant perceptual and memory systems which represent the speaker’s external and internal world. The form-generating system has access to a
mental syllabary. Let us now turn to the processing
components in slightly more detail.
CONCEPTUAL PREPARATION
PETER INDEFREY a n d WILLEM J. M. LEVELT
M a x Planck
Institute for Psycholinguistics, Nijmegen, The Netherlands
In preparing a message
for expression, we exercise our social and rhetorical
competence. An effective utterance will mind the knowledge state of the listener, the intention to be realized, the
845
59.1 Framework of processing components involved
in speech production. (From Levelt, 1999.)
FIGURE
achieved state of discourse, the attentional focus of the
interlocutor, and so on (Clark, 1996). Conceptual preparation capitalizes on our “theory of mind” skills—the
ability to estimate an interlocutor’s state of relevant beliefs and desires (Premack and Premack, 1995). All this is
subsumed under macroplanning (Butterworth, 1980; Levelt, 1989). One important aspect of macroplanning is
“linearization”—deciding what to say first, what to say
next, etc. (Levelt, 1981). This involves both rhetorical
decisions about how to guide the listener’s attention and
efficient management of working memory.
There is, in addition, microplanning. To be expressible
in language, a conceptual structure must be in a special,
“propositional” format. Visual images, musical patterns,
and motor images are typically in a different representational format. If they are to be expressed linguistically,
they must be recoded. This recoding is flexible, and dependent on the communicative goals. The same visual
image of a sheep and a goat juxtaposed can be expressed as “there is a sheep, and a goat to the right of it”
or “there is a goat, and a sheep to the left of it” (and in
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LANGUAGE
many other ways). This phenomenon, called perspective
taking, is not limited to the recoding of visual representations (cf. Levelt, 1996; Clark, 1997). The terminal elements in the propositional format must be lexical concepts,
concepts for which there are words in the language. The
choice of lexical concept is an important aspect of perspective taking. There are always multiple ways to refer
to the same entity: The animal/dog/labrador frightened me
or the interval/consonant/fifth is out of place here. Perspective
taking is ubiquitous in language production. As speakers, we are continuously mediating between visual, motor, person, etc., imagery systems and semantic systems
of lexical concepts. This mediation is under the pressure
of communicative effectiveness. It wouldn’t be surprising if conceptual preparation turns out to be a widely
distributed cerebral affair.
ENCODING The lexical concepts that
are activated in constructing a message for expression
trigger the retrieval of lemmas from the mental lexicon.
These are syntactic words, characterized by a syntactic
GRAMMATICAL
frame. There is a lemma for each lexical concept and for
all function words. Syntactic word frames specify, among
other things, how semantic arguments in the message
(such as theme or recipient) should be mapped onto syntactic functions (such as direct or indirect object). In Sally
gave Peter a bike, the recipient of Sally’s giving is Peter and
the theme is a bike. The syntactic frame of give moves the
corresponding lemmas into indirect and direct object position, respectively. The syntactic frames of selected lemmas (verbs, nouns, etc.) combine and recombine to build
a syntactic pattern for the message as a whole, a “surface
structure.” Surface structures are incrementally created. As
soon as a first lemma is selected, syntactic construction is
initiated, and it keeps going as further lemmas become
available. These processes are typically disturbed in
agrammatic patients.
A first major step
in the generation of the articulatory shape of an utterance involves the creation of phonological words and
phrases and the generation of intonational phrases. A
core process here is the retrieval of phonological codes.
Once selected, a lemma activates the phonological
codes of each of its morphemes. For instance, after selection of the noun lemma postbox, the codes for each of its
morphemes post and box are activated: /pE*st/, /bAks/.
Most neuroimaging work in word production involves
monomorphemic words and hence reveals nothing
about the production of complex morphology.
Accessing a word’s or morpheme’s phonological code
is no trivial matter, neuropsychologically speaking.
Anomic disorders, for instance, are often blockades of
phonological access with preserved access to syntactic information. Badecker, Miozzo, and Zanuttini (1995), for instance, reported the case study of an Italian anomic
patient who is unable to name any picture, but in all cases
knows the gender of the target word. Gender is a syntactic
word property, encoded in the lemma. Jescheniak and
Levelt (1994) have shown that the “word-frequency effect” (i.e., picture naming is slower when the name is a
low-frequency word than when it is a high-frequency
word) emerges during the retrieval of a word’s phonological code. It does not arise at the level of lemma selection.
Clearly, there is a dedicated system involved in the storage and retrieval of phonological codes.
The primary use of phonological codes is the generation of syllabic structure. The domain of syllabification is the phonological word. Syllabification doesn’t
respect lexical boundaries. In the phrase I understand it,
the syllabification becomes I un-der-stan-dit, where the
last syllable (dit) straddles a lexical boundary; understandit is a single phonological word. Syllabification also
depends on inflection—un-der-stand, un-der-stands, un-derMORPHOPHONOLOGICAL ENCODING
stan-ding—it is a highly context-dependent process. Most
probably, a word’s syllabification is not stored in its
phonological code. The incremental syllabification of
phonological words in connected speech is an independent computational process (cf. Levelt, Roelofs, and
Meyer, 1999, for a detailed theory of phonological
word formation).
As the surface structure expands, the speaker also
composes larger phonological units. One such unit is the
phonological phrase. It is a metrical unit. It tends to start
right after the lexical head of a surface phrase (i.e., right
after the noun of a noun phrase, or right after the main
verb in a verb phrase), and it leads up to include the
next lexical head. Here is such a metrical grouping: the
fellow/that I sought/was standing/near the table/. Within a
phonological phrase, there is so-called nuclear stress on
the lexical head word.
Phonological phrases combine into smaller or larger
intonational phrases. These are sense units that are characterized by their intonation contour. The whole of the example sentence above can be cast as a single intonational
phrase. Pitch movement will then lead up to the nuclear
tone, which consists of a pitch accent on the first syllable
of table (ta-), followed by a boundary tone on the last syllable of the phrase (-ble). Falling boundary tones suggest
completion, whereas rising boundary tones invite continuation either on the part of the speaker or on the part of
the interlocutor. The ultimate output of morphophonological encoding is called the phonological score (in analogy
to a musical score).
ENCODING The incremental generation of
metrically grouped and pitch-marked phonological syllables is closely followed by the generation of gestural
patterns for these syllables in their larger context. It is
largely unknown what kind of processing mechanism
creates these gestural scores. The system must be generative in that speakers can produce syllables that they
never produced before (in reading nonsense words for
instance). Still, it appears from language statistics that
speakers of English or Dutch do some 85% of their
speaking with no more than 500 different syllables (out
of more than 12,000 different syllables; cf. Schiller et al.,
1996). In these languages, speakers hardly ever produce
an entirely new syllable. Also, many languages (such as
Mandarin Chinese) have no more than a few hundred
different syllables. Hence, it is reasonable to assume that
these highly overused articulatory routines are stored
somewhere in the brain, and the premotor cortex is a
good candidate (cf. Rizzolatti and Gentilucci, 1988).
This repository of gestural scores is called the mental syllabary (Levelt, 1992). The generated gestural pattern for
an utterance is called the articulatory or gestural score.
PHONETIC
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
ARTICULATION Whatever the origin of the articulatory
score, it is ultimately executed by the laryngeal and supralaryngeal systems. These are under the control of the
larynx and face area of the somatosensory cortex, caudal midbrain structures, and cerebellum. Articulatory
execution is quite flexible. The same articulatory target
can often be realized in different ways. The system tends
to minimize effort, given the prevailing physical contingencies. It is, for instance, possible to speak intelligibly
with food or even a pipe in the mouth. Articulation is
our most sophisticated motor system. It is normal to produce some 12 speech sounds (consonants, vowels) per
second, and this involves control over some 100 different muscles. This masterpiece is achieved by concurrent, overlapping execution of articulatory gestures
(Liberman, 1996).
SELF PERCEPTION, MONITORING, A N D REPAIR
Speak-
ers are their own listeners. Whether listening to one’s
own speech or listening to somebody else’s speech, the
same superior temporal lobe structures are activated
(McGuire, Silbersweig, and Frith, 1996, Price, Wise, et
al., 1996). This feedback is one way for the speaker to
exercise some degree of output control. For instance, we
immediately adapt the loudness of our speech to the
prevailing noise in our speech environment. We also
tend to correct obvious or disturbing output errors or
infelicities. This self-monitoring, however, is not based
solely on the feedback of overt speech. We can also
monitor our internal speech and catch an error before
the word is (fully) pronounced (as in: we can go straight to
the ye-, to the orange node, where the almost-error here is
yellow). What is this internal speech? As Wheeldon and
Levelt (1995) have experimentally argued, it’s likely
that what we monitor for in internal speech is the phonological score, i.e., the output of morphophonological
encoding.
This bird’s eye view of the speaker’s functional organization provides us with the further layout of this chapter. We first discuss the many neuroimaging studies in
word production. In that discussion, we are guided by a
stage theory of word production, as diagrammed in figure 59.1. Following that, we turn to the few studies of
grammatical encoding and to some studies of internal
speech and self-monitoring.
Producing words: A task analysis
In neuroimaging studies of word production, we encounter a rich variety of tasks—verb generation, noun
generation, picture naming, word reading, word repetition, generating words starting with a particular letter,
and the like. The choice of experimental tasks and con848
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trols demonstrates both inventiveness and ingenuity, but
may also carry with it some degree of arbitrariness. Subtraction studies, in particular, are based on a difference
logic that requires a componential analysis of the functional organization involved in the experimental and
control tasks. It is rare, however, that such a componential analysis is independently performed and tested—say,
by way of reaction time studies. Pending such task analyses, the present review can provide only a theoretical
handle, presenting a componential analysis of normal
word production based on the theoretical framework in
figure 59.1. A fuller, comprehensive account of that
functional word production theory can be found in Levelt, Roelofs, and Meyer (1999).
The left panel of figure 59.2 represents the components—the “core processes”—involved in word production, as derived from figure 59.1. As far as word
production is concerned, the core aspect of conceptual
preparation is to map some state of affairs onto a lexical
concept. The state of affairs can be a perceptual image
(as in picture naming), the image of an activity (as in verb
generation), and so forth. In all cases we find perspective
taking—a decision on the type of lexical concept that is
apparently wanted in the experimental task. (For instance, one must decide whether to name an object by its
basic level term, such as dog in normal picture naming, or
to use a superordinate term, such as animal in a semantic
categorization task.) The grammatical encoding aspect of
word production is lemma access—selecting the appropriate syntactic word. It is at this step that the word’s syntactic properties, such as gender, mass/count noun,
syntactic argument structure, etc., become available.
There are two major aspects to morphophonological encoding, now distinguished in figure 59.2. The first one,
morphological encoding, provides access to the word’s
morphological structure and the phonological codes of
each morpheme. For the monomorphemic words used
in almost all neuroimaging studies, this stage is just accessing the word’s phonological code. An important independent variable here, affecting just this stage, is
word/morpheme frequency. The second one, phonological encoding proper, is the incremental construction of
the phonological word and in particular the word’s syllabification in context. This is probably the word representation figuring in internal speech. It may be (but need
not be) the end stage in silent word generation tasks. The
next component, phonetic encoding, provides a gestural
or articulatory score for the word. It is likely that highly
practiced syllabic motor routines are accessed at this
stage. In the final stage of word production, the constructed or retrieved gestural score is executed by the articulatory apparatus, resulting in an overt acoustic signal,
the spoken word. In all nonsilent word generation tasks
FIGURE 59.2 Core processing stages in the production of
words and the involvement of core and lead-in processes in
various word production tasks. A check mark indicates involvement of the component process in the task. A check mark
in parentheses indicates that the component’s involvement depends on details of the task. Phonetic encoding and articulation, for instance, are involved in overt, but not in silent word
production tasks.
there is auditory feedback, triggering the speaker’s normal word perception system. But there is feedback in silent word generation too, probably from the level of
phonological word encoding.
The subsequent columns of figure 59.2 present a tentative analysis of the various word production tasks reviewed in this chapter. In particular, these columns mark
the core processing components that are probably involved in these tasks. This aspect of the task analysis is
relatively straightforward (though not at all inviolable).
Much more problematic is the analysis of what we call
the task’s “lead-in.” Different tasks enter the componential structure depicted in the left panel at different levels.
In picture naming, for instance, the task enters the componential hierarchy from the very top component, conceptual preparation. The lead-in process is visual object
recognition, which provides an object percept as input
to conceptual preparation. Compare this to pseudoword
reading. Here the hierarchy is probably entered at the
level of phonological encoding—there is no accessing of
a syntactic word or of a word’s phonological code, but
there is syllabification. The lead-in process is visual
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
orthographic analysis, some kind of bottom-up grapheme-to-phoneme mapping, which provides the ordered
pattern of phonemes as input to syllabification. These
lead-in processes are the real bottleneck for neuroimaging studies in word production. They are usually easily
invented but ill-understood; still, they always contribute
essentially to the neuroimaging results. Without serious
behavioral research, one can only speculate at the processes involved in most task lead-ins. The top row of the
columns in figure 59.2 provides some hunches about the
lead-in processes involved in the various neuroimaging
studies of word production. Here, we discuss just seven
of them.
Picture naming Here the lead-in process is visual object
recognition. It is the best understood lead-in process.
Still, many variables are to be controlled, including visual complexity, perspectival orientation of the object,
color versus black-and-white, and, of course, object category. All core components of word production are involved in picture naming.
Ve r b g e n e r a t i o n This task also involves all core components of word production, but the lead-in process is illunderstood (cf. Indefrey, 1997). The subject sees or
hears a noun, which triggers a visual or auditory word
recognition process. If the noun is a concrete one, the
subject will probably generate a visual image; and, under the perspective of the task, that image activates one
or more associated actions in long-term memory. These,
then, guide the further conceptual preparation. When
the noun is abstract, long-term memory may be accessed without visual imagery. But there are possible
shortcuts, too. A perceived noun may directly activate a
verbal concept or even a verb lemma by sheer association, as in knife–cut.
Noun generation The typical task here is to present a semantic category, such as “jobs” or “tools” or “animals,”
and the subject is asked to generate as many exemplars
as possible. It is a so-called “word fluency” task. The
lead-in process may involve something as complicated
as an imaginary tour, such as mentally touring a zoo, or
it may be a much lower-level process, such as word association. And the subject’s strategy may differ rather drastically for different semantic categories. But it is quite
likely that, at least from lexical selection on, all core processes of word generation are involved.
Generating words from beginning letter(s) The lead-in process is quite enigmatic. The letter “a” is a preferred stimulus. Like most other letters, it does not represent a
unique phoneme in English, and the task probably capi850
LANGUAGE
talizes on visual word imagery. We can apparently retrieve orthographic word patterns beginning with “a.”
The same holds for so-called “stem completion”—transforming a word-beginning like “gre-” to its completed
form “green.” These visually imaged patterns are then
“read,” occasionally involving some semantic activation.
From there on, we are back to the core process, somewhere beginning at morphological or phonological
encoding.
Word repetition The subject repeats a heard word. The
lead-in process involves auditory word recognition, at
least to some extent. We can repeat words we don’t understand (and nonwords for that matter); hence it suffices to have the phonological parse of the word. From
there, the core process can be triggered at the level of
phonological encoding (we must syllabify the word).
Still, the lead-in process may be a lot richer, involving
activation of the full lexical concept. In delayed word
repetition tasks, an “articulatory loop” is involved—the
subject rehearses the word during the delay.
Word reading The lead-in process is visual word recognition, which is complicated enough by itself. The core
process may start at the low level of phonological encoding, from the set of activated phonemes (the phonological route); or it may start all the way up from the
activated lexical concept (the semantic route). The strategy may differ from subject to subject, even from word
to word.
Pseudoword reading Here, only the phonological route is
available after the visual lead-in process. Although nonwords can have morphology (as in “Jabberwocky”), that
was never the case in the tasks reviewed here. Hence,
phonological encoding is the first core process in a
pseudoword reading task.
In the following we will, to the best of our abilities, acknowledge the components involved in both the experimental word production tasks and their controls. But
given the present state of the art, this is not always possible.
Cerebral localizations for word production—
A meta-analysis
Research on brain regions involved in word production
has been carried out with a wide variety of techniques.
Among these are the study of brain lesions, direct cortical electrical stimulation, cortical stimulation by means
of implanted subdural electrode grids, recording and
source localization of event-related electrical and magnetic cortical activity (ERP, MEG, subdural electrode
grids, single-cell recordings), and measurement of re-
gional cerebral metabolic and blood flow changes (PET,
fMRI). Clearly, these techniques have contributed to
our present knowledge on the neural substrates of single
word production in different ways. Take cortical stimulation for example. Usually applied in the context of impending surgical interventions, cortical stimulation has
provided evidence on loci which, when temporarily inactivated, impair word production—i.e., loci that are in
some way necessary for the production process. But this
technique is applied only to locations where languagerelated sites are suspected and then only to the limited
part of the cortex that is exposed. In contrast, PET and
fMRI can, in principle, reveal all areas that are more
strongly activated during word production—including areas that may not be essential to the process and/or those
whose impairment leads to no detectable difference in
performance. ERP and MEG have provided preliminary insights in the temporal course of cortical activations. The sources of event-related electrical or magnetic
activations can be localized. There are, however, limitations inherent in the mathematical procedures involved,
so that, at present, the spatial information provided by
these methods is considered less reliable. Due to the nature of the signal, subcortical structures are largely invisible to electrophysiological methods.
The purpose of this section is to combine the evidence provided by all these techniques and to give an
overview of the localization (and to some extent the
temporal order) of cerebral activations during word
production. Furthermore, we will try to identify the
neural substrates of the different processing components laid out in the previous section. To this end, we
analyze the data reported in a large number of studies
according to the following heuristic principle: If, for a
given processing component, there are subserving
brain regions, then these regions should be found active in all experimental tasks sharing the processing
component, whatever other processing components
these tasks may comprise. In addition, the region(s)
should not be active in experimental tasks that do not
share the component.
This approach allows for the isolation of processing
components between studies even if isolation within single studies is not possible due to the difficulty in controlling for lead-in processes. Nevertheless, four conditions
must be met. First, the processing components must be
independently defined, so that their absence or presence
can be evaluated for every experiment by applying the
same criteria (which may differ from the author’s criteria). Second, the task and control conditions must be
heterogeneous enough across different experiments to
ensure that a specific processing component is the only
shared component. Third, the task and control condi-
tions must be heterogeneous enough across different experiments to ensure that for every processing
component there is a different set of tasks that share the
component. Fourth, the data base must be large, comprising enough experiments for a reliable identification
of activations typically found for the different tasks. For
word production these requirements seem to be sufficiently met, considering that word production tasks
have been among the most frequently applied language
tasks in neurocognitive research.
We analyzed the cerebral localization
data from 58 word production experiments (table 59.1).
Our focus was on the core process of word production;
thus, we excluded experiments reporting enhanced cerebral activations during word production tasks relative
to control tasks that comprised most or all of the word
production process—reading aloud, for example (Petersen et al., 1989; Raichle et al., 1994; Buckner, Raichle, and Petersen, 1995; Snyder et al., 1995; Fiez et al.,
1996; Abdullaev and Posner, 1997), or object naming
(Martin et al., 1995). Nor did our approach allow for inclusion of experiments or task comparisons focusing on
the relative strengths of components of the word production process—comparisons of reading regularly
spelled versus irregularly spelled words, for example
(Herbster et al., 1997). Activations of these two tasks relative to baseline, however, were included. It was assumed throughout that the reported activation foci
reflected true increases during the tasks rather than decreases during the baseline conditions.
Combining data from different techniques made it
necessary to find a common term for cerebral localizations observed in relation to certain tasks. Since the majority of experiments employed P E T or fMRI, we will
use the terms “activations” or “activated areas,” extending that usage to E E G and M E G sources and to sites
where cortical stimulation or lesions interfere with certain functions. We are aware that, for the latter case, one
can at best infer that such locations are “active” in normal functioning.
The double reference system for anatomical localizations adopted here was used in order to capture data
on the localization of cerebral activations stemming
from methods with different resolution. On a gross
level, comparable to a high degree of filtering, the reported loci were coded in a descriptive reference system dividing the cerebral lobes into two or three
rostrocaudal or mediolateral segments of roughly equal
size. Activations of cingulate, insula, and cerebellum
were only differentiated in left and right. The segment
labels were defined in terms of Talairach coordinates as
given in table 59.2 (top).
PROCEDURE
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
TABLE 59.1
Overview of experiments included in the word production data set
Task
Picture naming
aloud
Authors, methods, control conditions
Ojemann (1983)
Cortical/thalamic
stimulation
Haglund et al. (1994)
Cortical stimulation
Ojemann et al. (1989)
Cortical stimulation
Schäffler et al. (1993)
Cortical stimulation
Crone et al. (1994)
Subdural grid
Salmelin et al. (1994)
MEG
Bookheimer et al. (1995)
PET, nonsense drawings
Damasio et al. (1996)
Lesion data
Abdullaev & Melnichuk
(1995)
Single-cell recordings,
blank screen
Damasio et al. (1996)
PET, “faces,” up/down
Levelt et al. (1998)
MEG
Picture naming
silent
Bookheimer et al. (1995)
PET, nonsense drawings
Martin et al. (1996)
PET, nonsense objects
Price, Moore, et al. (1996)
PET, objects, “yes”
Word generation
silent, verbs
Wise et al. (1991)
PET, rest
Crivello et al. (1995)
PET, rest
Poline et al. (1996)
PET, rest
Word generation
silent, nouns
Warburton et al. (1996)
PET, rest
Paulesu et al. (1997)
fMRI, rest
Generation from
initial letter(s)
Aloud:
Buckner, Raichle, &
Petersen (1995)
PET, silent fixation
Word reading
aloud
Ojemann (1983)
Cortical stimulation
Price et al. (1994)
PET, false fonts,
“ab-/present”
Gordon et al. (1997)
Cortical stimulation
Howard et al. (1992)
PET, false fonts, “crime”
Bookheimer et al. (1995)
PET, nonsense drawings
Petersen et al. (1989)
PET, fixation
Petersen et al. (1990)
PET, fixation
Price, Moore, &
Frackowiak (1996)
PET, rest
Beauregard et al. (1997)
Concrete, abstract,
emotional words (3)
PET, word reading
instructions + fixation
Pseudoword
reading aloud
Sakurai et al. (1993)
PET, fixation
Indefrey et al. (1996)
PET, false font strings
Herbster et al. (1997)
PET, letter strings, “hiya”
Pseudoword
reading silent
Petersen et al. (1990)
PET, fixation
Fujimaki et al. (1996)
MEG
Hagoort et al. (1999)
PET, fixation
Word repetition
aloud
Petersen et al. (1989)
PET, silent listening
Howard et al. (1992)
PET, reversed words,
“crime”
Gordon et al. (1997)
Cortical stimulation
Word reading
silent
Price et al. (1996b)
PET, rest
Pseudoword
repetition
silent
852
Warburton et al. (1996)
PET, rest
LANGUAGE
Warburton et al. (1996)
Exp. 1B, 2B+C, 3A (4)
PET, rest
Silent:
Paulesu et al. (1997)
fMRI, rest
Sakurai et al. (1992)
PET, fixation
Price, Moore, &
Frackowiak (1996)
PET, rest
Herbster et al. (1997)
Regular and irregular
words (2)
PET, letter strings, “hiya”
Sakurai et al. (1993)
PET, fixation
Rumsey et al. (1997)
PET, fixation
Bookheimer et al. (1995)
PET, nonsense drawings
Menard et al. (1996)
PET, xxXxx
Hagoort et al. (1999)
PET, fixation
Rumsey et al. (1997)
PET, fixation
Crone et al. (1994)
Subdural grid
More detailed anatomical references were additionally coded on a finer level in terms of gyri and subcortical structures following Talairach and Tournoux (1988).
At this level, cingulate, insular, and cerebellar activations were further differentiated descriptively (table
59.2, bottom). In this way, it was possible to capture the
fact that a PET activation focus reported as, for example, left inferior temporal gyrus, BA 37, would be consistent with electrophysiological data reporting a posterior
temporal source localization or with patient data reporting a left posterior temporal lesion. Note that the sum of
activations on the detailed level does not equal the number of activations on the gross level. A location, for example, that was reported as posterior temporal would be
marked only on the gross level. Conversely, two posterior temporal locations in the superior and middle temporal gyrus were marked as such on the detailed level
but only once on the gross level.
The studies included in this meta-analysis were not
given any weights reflecting reliability differences due
to design or size. This means that a certain degree of
overlap of activations between studies was considered
meaningful, but should not be interpreted as statistically
significant. Nonetheless, the notion of “meaningfulness”
was not totally arbitrary, but based on the following
quasi-statistical estimate: At the gross level of description, there were on average 6.5 activation sites reported
per experiment. Given that on this level of description
there were 28 regions of interest, any particular region
had a chance of less than one-fourth to be reported as
activated if reports were randomly distributed over regions. At the finer level of description, the average
number of reported activation sites per experiment was
8.8 and there were 104 regions of interest; thus each
had a chance of less than one-tenth to be reported as activated. Assuming these probabilities, the chance level
for a region to be reported as activated in a number of
studies was given by a binomial distribution. We rejected the possibility that the agreement of reports
about a certain region was coincidental if the chance
level was less than 10%. At the finer level of description,
this corresponded to minimally two reported activations for regions covered by less than six experiments,
minimally three reported activations for regions covered by six to eleven experiments, and so forth (4 out of
12–18; 5 out of 19–25; 6 out of 26–32). Note that for regions covered by many experiments a relatively smaller
number of positive reports was required to be above
chance (comparable to the fact that getting a 6 five
times with ten dice throws is less likely than getting one
6 with two dice throws). But this criterion does not
mean that atypical findings of activations in any single
study are necessarily coincidental. In most cases, the
number of experiments not reporting activations was
not sufficient to consider a region as inactive at the chosen error probability level. Rare observations do not,
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
therefore, exclude the possibility that a region is active.
They may, for example, reflect smaller activations that
are only detectable with refined techniques or better
scanning devices.
A second, related point is that the nature of the data
does not allow for an interpretation in terms of relative
strengths of activations of certain areas. It is known that
parameters such as item duration and frequency
strongly influence the resulting pattern of activations
(Price et al., 1994; Price, Moore, and Frackowiak, 1996).
It is thus possible that areas are more frequently found
active in some tasks, because their “typical” item durations and frequencies are higher or lower than in other
tasks. It seems wise not to overinterpret the data, given
that there is a considerable variability of these parameters across the studies of our data base; also, the interactions of these parameters with other experimental
factors are largely unknown.
PROCESSES According to our task analysis,
nonshared activations in picture naming and word generation (table 59.3, first two columns) cannot be related
to the core process of word production. The two tasks
differ not only with respect to their lead-in processes but
also with respect to the processes of phonetic encoding
and articulation, given that word generation was performed silently in all experiments of the data set,
whereas the majority of picture naming experiments
involved overt articulation with silent control tasks (see
table 59.1). Hence, to study the lead-in processes of picture naming, we must take into account activations that
were specific for picture naming when compared to
word generation and at the same time were not specific
for overt (in contrast to silent) naming in general (table
59.3, last two columns). Although such lead-in activations were fairly numerous, we cover only the two most
conspicuous sets here.
Six regions were reported as activated during word
generation but rarely activated during picture naming:
the left anterior superior frontal gyrus, the right anterior
insula, the right mid superior and middle temporal gyri,
the left caudate nucleus, and the right thalamus. While
activations of the left anterior frontal and middle temporal gyri, right anterior insula, and left caudate seem to be
specifically related to lead-in processes of word generation (see also Fiez, Raichle, and Petersen, 1996), the
case is different for the right mid superior temporal
gyrus and the right thalamus, which are also found in
word reading.
Te n regions were activated during picture naming but
not or only rarely during word generation: the left anterior insula, the left posterior inferior temporal and fusiform gyri, the medial occipital lobe bilaterally, the right
LEAD-IN
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LANGUAGE
caudate nucleus, the left midbrain, and the medial and
right lateral cerebellum (see, however, Fiez and Raichle,
1997, for evidence on right cerebellar activations in word
generation when directly compared with word reading or
picture naming). Five of these—the left anterior insula, the
left posterior fusiform gyrus, the left and right medial occipital lobe, and the right medial cerebellum—were also
found active in word reading, suggesting an involvement
in visual processing, the principal lead-in component of
picture naming and reading. Taking into account that activations of the posterior fusiform gyrus or the insula were
not reported for pseudoword reading, it seems that these
two areas may play a role at later visual processing stages,
such as the retrieval of visual word forms or object patterns (cf. Sergent, Ohta, and MacDonald, 1992). In contrast, medial occipital activations observed during word
and pseudoword reading are demonstrably due to the
processing of nonlinguistic visual features of word-like
stimuli, such as string length (Beauregard et al., 1997;
Indefrey et al., 1997).
T H E C O R E PROCESS OF W O R D PRODUCTION
Accord-
ing to our task analysis, picture naming and word generation are the two tasks that include all components of
word production. The set of regions reported as activated
for both tasks (table 59.3, first two columns) can be considered as being related to the core process of word production up to and including phonological encoding. This
word production network is strictly left-lateralized, and
consists of the posterior inferior frontal gyrus (Broca’s
area), the mid superior and middle temporal gyri, the posterior superior and middle temporal gyri (Wernicke's
area), and the left thalamus.
By taking into account further tasks that enter the
word production process at different stages, we now attempt to identify the subprocesses to which these regions are particularly sensitive.
Conceptual preparation and lexical selection The activation
of a lexical concept and the subsequent selection of the
corresponding lemma are processes that are shared by picture naming and word generation but not necessarily by
word reading. The synopsis of reported results yielded one
area within the word production network—the mid
segment of the left middle temporal gyrus—that has been
found activated during picture naming and word generation, but less so during word reading. This area is the best
candidate for a neural correlate of conceptual and/or lexical selection processes in word production. The mid part
of the left middle temporal gyrus has been found as part
of a “common semantic system” in a study (Vandenberghe
et al., 1996) involving word and object stimuli. This finding is compatible with a role of this region in conceptual
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
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LANGUAGE
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
*The number of activations reported for a region is given in proportion to the number of studies covering it. Relative cell frequencies exceeding the error probability threshold of p < .1 are printed in bold. Data were collapsed with respect to overt versus
silent responses in the last two columns. Aloud tasks with aloud control conditions are grouped with silent tasks.
Key: Except for SMA (= supplementary motor area), the abbreviations of gyri and subcortical structures follow Talairach and
Tournoux (1988): GFs, GFm, GFi = superior, middle, and inferior frontal gyrus; GFd = medial frontal gyrus; GO = orbital gyri;
GR = gyrus rectus; Gs = gyrus subcallosus; GPrC = precentral gyrus; GTs, GTm, GTi = superior, middle, and inferior temporal
gyrus; GF = fusiform gyrus; Gh = parahippocampal gyrus; GL = lingual gyrus; GpoC = postcentral gyrus; LPs, LPi = superior
and inferior parietal lobule; PCu = precuneus; Gsm = supramarginal gyrus; Ga = angular gyrus; Sca = calcarine sulcus; Cu =
cuneus; GOs, GOm, GOi = superior, middle, and inferior occipital gyri; NL = lenticular nucleus.
processing. It should, however, be kept in mind that the
activation of a lexical concept for word production is only
one very specific conceptual process among many other
conceptual-semantic processes. It is, more precisely, to be
distinguished from the semantically guided search processes in word generation (possibly subserved by anterior
frontal regions), as well as from prelinguistic conceptual
processes involved in object recognition and categorization [possibly subserved by the ventral temporal lobe and
a heterogeneous set of category-specific regions (cf. Martin et al., 1995, 1996; Damasio et al., 1996; Beauregard et
al., 1997)]. As far as the core process of word production
is concerned, these conceptual processes are to be considered as lead-in processes.
Phonological code retrieval Lexical word form retrieval
takes place in picture naming, word generation, and word
reading, but not in pseudoword reading. This pattern is
found in the reports on activations of the left posterior superior and middle temporal gyri, i.e., Wernicke’s area,
and the left thalamus. The posterior superior temporal
lobe has also been found active during word comprehension (Price, Wise, et al., 1996). It is thus conceivable that
a common store of lexical word form representations is
accessed in word production and comprehension.
Phonological encoding All tasks, including word repetition and pseudoword reading, involve the production of
phonological words. Neural structures subserving this
process should consequently be found active throughout. No region fulfills this requirement perfectly, but the
left posterior inferior frontal gyrus (Broca’s area) and the
left mid superior temporal gyrus come very close. Both
regions just miss our criterion for a meaningful number
of activations in tasks for which there are only few studies in the data set (word repetition, word generation
from initial letters). Cabeza and Nyberg (1997) reviewed
one repetition and two reading aloud experiments with
Broca’s area as the only active region in common. According to our task analysis, the only common processing component was indeed phonological encoding.
Broca’s area has been observed to be active not only
during explicit but also during implicit processing of
pseudowords (performing a feature detection task; Frith
et al., 1995; Price, Wise, and Frackowiak, 1996). The left
mid superior temporal gyrus, however, was not found
active in these studies, which may indicate a functional
difference between the two areas within nonlexical phonological processing.
Broca’s area is, furthermore, known to be activated in
tasks involving phonological processing in language
comprehension (Démonet et al., 1992,1996; Zatorre et
al., 1992; Fiez and Raichle, 1997). The common denom-
inator of these observations and the activation of Broca’s
area in language production seems to be that this region
is a nonlexical phonological processor.
Price and Friston (1997) presented a statistical
method—conjunction analysis—to isolate common processing components between different experiments.
They used this method to identify a processing component, which they called phonological retrieval, across
four word production experiments. The following areas
were reported to be related to this processing component: the left posterior basal temporal lobe BA 37 (this
area corresponds to the fusiform gyrus in our terminology), the left frontal operculum, the left thalamus, and
the midline cerebellum. Given that “phonological retrieval” according to our analysis corresponds to the two
processing stages of phonological code retrieval and
phonological encoding, the observed activations of the
left thalamus and the left frontal operculum are in good
agreement with the results we obtained here on the basis
of a large number of reported experiments. The other
two areas according to our analysis subserve a high-level
visual lead-in process and the articulatory process. Price
and Friston (1997) assume these processes to be shared
by the control conditions (viewing objects or strings of
false fonts and saying “yes” to every stimulus), leaving
only “phonological retrieval” as the common component of all task-control contrasts. It should, however, be
noted that this assumption holds only if the contributions of these task components are constant in the active
task and the baseline task. The authors themselves point
out that this core assumption of the cognitive subtraction
paradigm is problematic. Visual and motor-related activations have been observed to be modulated (e.g., by attention or response selection) in active relative to
passive tasks (Friston et al., 1996; see also Shulman et al.,
1997, for medial cerebellar activations in controlled motor response tasks). It is therefore not excluded that activations related to visual and articulatory processing
were equally enhanced in all four active tasks and consequently not filtered out by the conjunction statistics.
Phonetic encoding and articulation Activations related
to the production of the abstract articulatory program
and its execution should be found in tasks with overt
pronunciation and silent control conditions, but not in
silent tasks or in tasks where articulation has been controlled for (table 59.3, last two columns). All aloud tasks
with silent controls led to activations of primary motor
and sensory areas, i.e., the right and left ventral (and, to
some extent, dorsal) precentral gyri and the right and
left ventral postcentral gyri; but in the group of silent or
controlled aloud tasks, such activations were rarely reported. This finding was expected, since these areas are
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
known to be involved in the sensorimotor aspects of articulation. Hence, it provides independent validation for
our analysis procedure. It can also be concluded that
output control conditions such as saying the same word
to every stimulus cancel out these sensorimotor activations effectively.
Further regions typically found in aloud but not in silent tasks were the left anterior superior temporal gyrus,
the right S M A , and the left and medial cerebellum. The
dissociation in cerebellar activity, with left and medial
parts being closely linked to motor output, confirms an
observation by Shulman and colleagues (1997); it is also
discussed in a comprehensive review of cerebellar activations by Fiez and Raichle (1997).
Our survey shows a complex pattern of reports on left
anterior temporal and S M A activations. Both regions
are related to overt pronunciation, but seem to be tasksensitive as well. While the temporal area is most frequently reported in overt picture naming, S M A activations (left and right) seem to be rare in this task. Also,
S M A activations, though more frequent for aloud tasks,
are observed to some extent in silent tasks as well. The
latter is not restricted to silent word production, but is
also found in tasks involving verbal working memory or
nonverbal imagination of movements (for a discussion,
see Fiez and Raichle, 1997). It may be concluded that
the S M A is in some complex way related to motor planning and imagination of articulation. Given that the instructions in what we have designated silent tasks ranged
from mere “viewing” to “thinking” to silent “mouthing”
of responses, it is not difficult to understand that the
S M A involvement may vary to a great extent between
tasks.
The word production network that has
been identified on the basis of a substantial number of
experiments makes sense. It is largely identical with the
set of regions found to be necessary for picture naming
in direct cortical stimulation studies. Given that these
comprised but a minority of the experiments analyzed
and furthermore concentrated on a single task, this is not
a trivial result. It means that the neuroimaging studies,
despite their heterogeneity of methods and tasks, captured the essential processes of word production. It also
means that the cerebral structures subserving these processes can be successfully distinguished from the large
number of cerebral activations related to task-specific
and experiment-specific processes by an appropriate
meta-analysis procedure. On the other hand, neither the
network as a whole nor any single region was found activated in all experiments. There are a number of reasons
for this. First, weaker activations may have been overlooked or not reported. In general, the statistical threshSUMMARY
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olds applied in neuroimaging experiments tend to be
conservative; moreover, some authors may have focused on robust findings, applying very strict statistical
thresholds that rendered minor activations insignificant.
There also was a tendency toward fewer activations in
older studies, where the technology could not reliably
detect as many activations. Second, although we focused
on experiments with low-level control conditions, so
that the word production process itself would not be obscured, we could not confidently eliminate this obscuring in all cases. Stereotype overt responses (for instance,
a “yes” response on all trials) may be retrieved from an
articulatory buffer, but may also be normally produced
(as is the case with meaningful response alternatives,
such as saying “up” or “down” depending on the orientation of the stimulus object in the baseline task)—
thereby taking away at least part of the activations due
to core word production processes. Considering these
points, it would have been misleading to apply the
above heuristic principle to every single experiment
rather than to sets of experiments using similar tasks, as
we have done here.
The time course of word production
Every
processing stage of word production takes time. Furthermore, as the above analysis suggests, cortical areas involved in word production are specialists for certain
processing components. Activations in these regions
should, therefore, have temporal properties that are
compatible with the durations of the different processing
stages. Table 59.4 summarizes the small number of studies that, to date, have provided timing information related to cortical areas involved in word production. We
compare these data with estimates for the processing
stages in picture naming given by Levelt and colleagues
(1998) based on work by Thorpe, Fize, and Marlot
(1996), Levelt and colleagues (1991), Roelofs (1997),
Wheeldon and Levelt (1995), and Va n Turennout, Hagoort, and Brown (1997, 1998).
It is estimated that visual and conceptual processing
are accomplished within the first 150 ms, and lexical
selection within 275 ms from picture onset. As
identified in the previous section, the corresponding
cortical sites were the medial occipital lobe (bilaterally),
the left medial posterior temporal lobe for visual processing, and the mid segment of the left middle temporal gyrus for conceptual preparation and lexical
selection. The time windows given for occipital activations in table 59.4 are in good agreement with what is
assumed for early visual processing. Inferior posterior
temporal activations during reading also seem to occur
T I M E W I N D O W S FOR C O M P O N E N T PROCESSES
TABLE 59.4
Time course of word production in relation to anatomical regions*
Picture naming
aloud
Task
Studies
Occipital
R
L
Parietal
R
Wo r d
reading
silent
Pseudoword
reading
silent
Wo r d
repetition
aloud
Crone et al.
(1994)
Salmelin et al.
(1994)
Levelt et al.
(1998)
Salmelin et
al. (1996)
Fujimaki et al.
(1996)
Crone et al.
(1994)
Subdural grid
MEG
MEG
MEG
MEG
Subdural grid
medial
0–200
0–275
lateral
0–275
medial
0–275
lateral
0–275
posterior
200–400 (Ga)
100–200
100–200
150–275 (Gsm)
>400 (cingulum)
anterior
>400 (cingulum)
sensory
L
400–600
posterior
200–400 (Ga)
anterior
sensory
400–600
Te m p o ra l posterior
R
100–200 (GTi)
mid
<400
300–500 (GTs)
anterior
L
posterior
mid
300–600
(GTs,Tm)
200–400 (GTs)
600–900 (GTs)
275–400 (GTs)
100–200 (GTi)
200–400
(GTs,GTm)
275–400 (GTs)
200–400
(GTs,GTm)
<400
300–900 (GTs)
anterior
Frontal
R
motor
500–600
posterior
200–400
200–400
anterior
L
motor
posterior
500–600
300–900 (GFi)
200–400
600–900 (GFi)
anterior
*Time intervals are given in milliseconds; additional anatomical information is given in parentheses. Ga = angular gyrus; Gsm =
supramarginal gyrus; GTs, GTm = superior, middle temporal gyrus; GFi = inferior frontal gyrus.
in this time window (Salmelin et al., 1996). Instead of
the expected middle temporal activation site, however,
both Salmelin and colleagues (1994) and Levelt and
colleagues (1998) identified dipoles in posterior parietal
regions, which have not been found active in PET
studies.
Lexical phonological code retrieval and phonological
encoding are estimated to take place between 275 and
INDEFREY AND LEVELT: LANGUAGE PRODUCTION
400 ms. The corresponding cortical sites identified
above were the left posterior superior and middle temporal gyri for accessing the form code, as well as the left
posterior inferior frontal gyrus and the left mid superior
temporal gyrus for phonological encoding. The time
windows given in table 59.4 clearly support an involvement of Wernicke’s area in word form retrieval. Further
support comes from a study by Kuriki and colleagues
(1996) reporting a time window of 210–410 ms in this region during a phonological matching task on syllabograms. The situation is less clear for phonological
encoding, where both supporting and disagreeing time
windows have been found for Broca’s area and the mid
segment of the left superior temporal lobe. The available
data on the time course of word production thus support
the localization studies with respect to lead-in processes
and word form access. They are consistent with respect
to phonological encoding, but raise doubt with respect
to the cortical sites related to conceptual preparation
and lemma access in word production.
INTEGRATING
SPATIAL
AND
TEMPORAL
INFORMA-
Taking into account both the results of the previous section and the present data on the time course of
word production, the following (tentative) picture of the
spatial and temporal flow of activation in word production has emerged: In picture naming and probably also
in word generation from visual stimuli (see Abdullaev
and Posner, 1997) visual and conceptual lead-in processes involving occipital, ventrotemporal, and anterior
frontal regions converge within 275 ms from stimulus
onset on a lexical concept to be expressed. In addition,
the best-fitting lexical item is selected within this period.
The middle part of the left middle temporal gyrus may
be involved in this conceptually driven lexical selection
process. Within the following 125 ms, the activation
spreads to Wernicke’s area, where the lexically stored
phonological code of the word is retrieved, and this information is relayed anteriorly to Broca’s area and/or
the left mid superior temporal lobe for post-lexical phonological encoding. Within another 200 ms, the resulting phonological word is phonetically encoded (with
possible contributions of S M A and cerebellum to this or
additional motor planning processes) and sensorimotor
areas involved in articulation become active.
TION
Syntactic production
Research on the neural correlates of syntactic processing has mainly concentrated on syntactic comprehension. Regions in and around Broca’s area have been
most frequently identified as being related to syntactic
processing (Stowe et al., 1994; Indefrey et al., 1996;
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LANGUAGE
Just et al., 1996; Stromswold et al., 1996). Caplan,
Hildebrandt, and Makris (1996), on the other hand, did
not find a significant difference in syntactic comprehension of agrammatic patients between anterior and
posterior sylvian lesion sites in a thorough study involving a range of syntactic constructions. Just and colleagues (1996), too, found Wernicke’s area (as well as
the right-sided homologs of Broca’s and Wernicke’s
area) to be sensitive to syntactic complexity. Tw o studies (Mazoyer et al., 1993; Dronkers et al., 1999) suggested a role for the left anterior superior temporal
lobe in syntactic processing. The pseudoword sentence
repetition task of Indefrey and colleagues (1996) comprised a syntactic production component in addition to
syntactic parsing, and the resulting activation focus was
more rostrodorsally located (border of Broca’s area
and the adjacent middle frontal gyrus, BA 9) than the
foci identified in pure comprehension tasks. Direct
electrical stimulation of a similar site was found by
Ojemann (1983) to interfere with the grammatically
correct repetition of sentences. In the latter study, however, a number of perisylvian stimulation sites yielded
the same effect, so that at present there is no real evidence for cortical areas specifically subserving syntactic production.
Self-monitoring
Self-monitoring (see figure 59.1) involves an external loop,
taking as input the acoustic speech signal of the
speaker’s own voice, and an internal loop, taking as input
the phonological score—i.e., the output of phonological
encoding. The most economical assumption is that both
loops enter the processing pathway that is used for normal speech comprehension (Levelt, 1989).
L O O P There is evidence that hearing
one’s own voice while speaking induces the same temporal lobe activations as listening to someone else’s
voice (McGuire, Silbersweig, and Frith, 1996; Price,
Wise, et al., 1996). McGuire, Silbersweig, and Frith, furthermore, were able to induce additional bilateral superior temporal activations by distorting the subjects’
feedback of their own voices or presenting the subjects
with alien feedback while they spoke. These results
show that just as in listening to other people’s speech
(Démonet et al., 1992; Zatorre et al., 1992) attentional
modulation of the activity of the temporal cortices in
self-monitoring is possible.
EXTERNAL
L O O P McGuire and colleagues (1996) also
provided some evidence that internal monitoring, too,
makes use of a cortical area involved in speech
INTERNAL
perception—more precisely, the left posterior superior
temporal lobe. This area showed stronger activation (together with motor areas) when subjects imagined hearing
another person’s voice than when they spoke silently to
themselves. It is not implausible that the observed blood
flow increase was due to an attentional modulation of internal self-monitoring, although other explanations are
possible as well.
Conclusion
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Speaking involves substantially more than merely producing words. The recent neuroscience literature on
speech production reviewed in this chapter is limited
owing to its emphasis on word production. But this may
well be a transient state of affairs. After all, the wealth of
data that enabled the present meta-analysis derives from
no more than a decade or so of neuroimaging research.
We have an emerging picture now of the cerebral network that underlies the production of words. A crucial
ingredient of this meta-analysis is the explicit, detailed
functional theory of word production. It provided us
with the framework for a componential task analysis and
with time frame estimates for the component processes.
No doubt, the same approach, applied to other aspects
of speaking, will eventually lead to similar progress.
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